A spawning aggregation of Nassau grouperEpinephelus striatus (Pisces: Serranidae) in the Mexican Caribbean

Synopsis An annual spawning aggregation of Nassau grouper,Epinephelus striatus, observed off the southern coast of Quintana Roo, Mexico, on full moon days in December and January, was surveyed during the winters from 1991 to 1993. A fish aggregation had been appearing at one traditional site off Mahahual for more than 80 years, but currently it forms at undetermined sites far from the original site. Fish migration was northwards in the fore reef border along the 14 m isobath and isolated groups were observed. Through visual assessment fish aggregations were found to include 200 to 500 individuals. The fishing on Nassau grouper aggregations has been active since 1910 by using hook and line; however, recently fishermen used gill nets. It is necessary to continue the research on this aggregation and establish management guidelines to protect the species.     

Spawning aggregations of three protogynous groupers in the southern Gulf of Mexico

Spawning aggregations of red hind Epinephelus guttatus, tiger grouper Mycteroperca tigris and yellowfin grouper Mycteroperca venenosa were identified at two coral‐reef systems: Arrecife Alacranes (emergent bank reef) and Bajos del Norte (submerged bank reef) on the continental shelf of the Yucatan Peninsula (Campeche Bank), Mexico. At both reefs, E. guttatus forms large spawning aggregations between February and March. At Bajos del Norte, M. tigris reproduces in a small, low‐density aggregation in May, while M. venenosa aggregates at high densities for spawning between March and May. Multi‐species use of an aggregation site by E. guttatus and M. venenosa was observed at Bajos del Norte. The identified spawning aggregations are apparently stable in location over time, and all three species were commonly observed to spawn within 1 week following the full moon. Development and survival of the larvae spawned in these aggregations are probably aided by a seasonal (spring–summer) upwelling in the north‐east Campeche Bank. A permanent area closure at Bajos del Norte, currently outside any specific fisheries management area or regulations, would provide protection needed for the spawning aggregations of these three species.     

Aggregations of <Emphasis Type="Italic">Plectropomus areolatus</Emphasis> and <Emphasis Type="Italic">Epinephelus fuscoguttatus</Emphasis> (groupers,Serranidae) in the Komodo National Park,Indonesia: Monitoring and Implications for Management

Synopsis We identify fishery management implications from a long-term monitoring program focusing on spawning aggregations of high valued reef fish in Komodo National Park (KNP), Eastern Indonesia. Management objectives of KNP are not only to protect biodiversity, but also to conserve spawning stocks of high-valued commercial species for the replenishment of surrounding fishing grounds. We monitored two sites twice monthly over five years for two species of grouper, Epinephelus fuscoguttatus and Plectropomus areolatus. One site had an aggregation of both E. fuscoguttatus and P. areolatus, whereas the other site contained an aggregation of P. areolatus only. Over the five years monitoring period, aggregations typically formed during each full moon between September and February. Additionally, P. areolatus occasionally aggregated during new moons between April and July. We observed spawning only once, but because formation of aggregations were correlated to a higher incidence of behavior and signs indicative of reproduction and because most fish present were adults, it is likely that the formation of aggregations was associated with spawning. Over the five years monitoring period there was a reduction in mean fish size of up to 8 cm for P. areolatus, and a reduction in numbers of aggregating E. fuscoguttatus. Despite limited protection initiated in 2001, both sites are still heavily fished by local artisanal fishers. Because the observed reductions in size and in numbers could be caused by fishing pressure, managers should follow the precautionary principle by putting additional protective management in place. Since both species are relatively long-lived, at least five years of continued monitoring may be necessary to determine the outcome of management intervention. The variability in timing of aggregation in respect to season and moon phase in P. areolatus indicates that long-term monitoring must cover the entire year and both moon phases.     

Observations of a Nassau grouper, Epinephelus striatus, Spawning Aggregation Site in Little Cayman, Cayman Islands, Including Multi-Species Spawning Information

Mass spawning aggregations of Caribbean grouper species are a conservation priority because of declines due to over-fishing. Previous studies have documented five historical aggregation sites in the Cayman Islands. Today, three of these sites are inactive or commercially extinct. In January 2002, the Reef Environmental Education Foundation led an expedition to Little Cayman Island to document a recently re-discovered spawning aggregation of Nassau grouper, Epinephelus striatus. A team of divers estimated the abundance, color phase composition, and courtship and spawning behavior of the aggregating grouper. The color phase composition of the aggregation shifted both during the course of each evening and throughout the 10-day project. Divers documented atypical coloration and courtship behavior in 10 additional fish species, of which five were seen spawning. Artisanal fishing occurred daily on the aggregation. The Cayman Islands Department of the Environment collected landings data and sampled catches to obtain length and sex ratios. The Cayman fishing fleet, while small, had a significant impact on the aggregation with a harvest of almost 2000 Nassau grouper during the 10-day project. The study site supports the largest known Nassau grouper aggregation in the Cayman Islands. The relatively large size of fish and the high proportion of males indicate that this site supports a relatively healthy aggregation compared to other Nassau grouper aggregation sites throughout the Caribbean.     

Preliminary observations of spawning behavior of white-streaked grouper (Epinephelus ongus) in an Okinawan coral reef

Epinephelus ongus forms spawning aggregations at specific reef sites at Okinawa. The aim of the present study was to describe the spawning behavior of E. ongus based upon underwater video imagery. Epinephelus ongus spawning is usually paired with courting males and females rushing upwards towards the surface during the spawning ascent, with a subsequent release of eggs at the apex of the ascent. In two of four spawning events we observed, however, the ascending pair was joined by additional males. Fish eggs collected at the site were identified as those of E. ongus and confirmed that this species spawns mainly at night during a slack tide. This study is the first documentation of E. ongus spawning behavior in the field.     

Historical and contemporary evidence of a mutton snapper (<Emphasis Type="Italic">Lutjanus analis</Emphasis> Cuvier, 1828) spawning aggregation fishery in decline

Scientific information on reef fish spawning aggregation fisheries is sparse in light of numerous regional declines and extirpations from overexploitation. Fisher interviews of the small-scale commercial mutton snapper (Lutjanus analis) spawning aggregation fishery at Gladden Spit, Belize, suggests a historic decadal decline. The reported trend is supported by analysis of inter-seasonal catch and effort and yield (2000–2002) that reveals a 59% decline in catch per unit effort (CPUE) and a 22% decrease in mean landings per boat. Declining population-level trends are also supported by a significant decrease in inter-annual median lengths of mutton snappers (2000–2006). These findings demonstrate the need for additional life history information that includes length-associated age and details on growth to provide clearer support of the effects on, and responses by, populations following fishing. In view of the historical changes to mutton snapper CPUE and landings at Gladden Spit and the fishery-associated declines in fish spawning aggregations observed globally, a precautionary approach to spawning aggregation management is warranted that provides full protection from fishing to enhance population persistence. The findings also highlight the need for substantially greater enforcement and long-term fisheries monitoring under a comprehensive regional management strategy.     

First record of a spawning aggregation for the tropical eastern Pacific endemic grouper Mycteroperca olfax in the Galapagos Marine Reserve

This study provides direct and indirect evidence of temporally and spatially consistent spawning aggregations for the grouper Mycteroperca olfax. Recently reported declines in population numbers, probably related to the direct targeting of aggregations by artisanal fishermen, highlight the urgent need for species‐specific management actions in the Galapagos Marine Reserve, such as minimum and maximum landing sizes, and the importance of protecting key aggregation sites with the declaration of no‐take areas and the establishment of total fishing bans during the reproductive season.     

Spawning aggregations of Lutjanus cyanopterus(Cuvier) on the Belize Barrier Reef over a 6 year period

Cubera snapper Lutjanus cyanopterus aggregated to spawn at Gladden Spit, a salient sub‐surface reef promontory seaward of the emergent reef and near the continental shelf edge of Belize. Their spawning aggregations typically formed 2 days before to 12 days after full moon from March to September 1998–2003 within a 45 000 m² reef area. Peak abundance of 4000 to 10 000 individuals was observed between April and July each year, while actual spawning was most frequently observed in May. Spawning was observed consistently from 40 min before to 10 min after sunset within a confined area ≤1000 m². Data suggested that cubera snapper consistently formed seasonal spawning aggregations in relation to location, photoperiod, water temperature and lunar cycle, and that spawning was cued by time of day but not tides. The cubera snapper aggregation site was included within the Gladden Spit Marine Reserve, a conditional no‐take fishing zone.     

Density‐dependent effects on initial growth of a branching coral under restoration

Coral reef restoration aims to help threatened coral ecosystems recover from recent severe declines. Here we address whether coral fragments should be out‐planted individually or in larger aggregations. Theory suggests alternative possible outcomes: whereas out‐plants within aggregations might suffer from heightened negative interactions with neighbors (e.g. competition for space), they may alternatively benefit from positive interactions with neighbors (e.g. buffering wave disturbances). On a degraded reef in the Caribbean (St. Croix, USVI), using out‐plants of the critically endangered staghorn coral Acropora cervicornis, we experimentally tested how aggregation density (1–20 out‐planted coral fragments spaced at approximately 5 cm) influenced initial coral growth (over 3 months). Coral growth declined as a function of aggregation size, and out‐plants within larger aggregations had fewer and shorter secondary branches on average, indicative of horizontal competition for space. Our results therefore suggest that wide spacing of individuals will maximize the initial growth of out‐planted branching corals.     

Spatial and temporal patterns of movement and migration at spawning aggregations of red hind, <Emphasis Type="Italic">Epinephelus guttatus</Emphasis>, in the U.S. Virgin Islands

This study compared the spatial and temporal patterns of red hind, Epinephelus guttatus, movement and migration from annual spawning aggregations on St Thomas (STT) and St Croix (STX), United States Virgin Islands. Around STT E. guttatus migrated 6–33 km from a functional spawning migration area of 500 km² and around STX E. guttatus migrated 5–18 km from an area of 90 km². Similarities between sites were found in regards to timing of movement, temporal and spatial changes in sex ratios, annual and lunar predictability and were synchronized with environmental cues. E. guttatus spawning aggregations in the Virgin Islands occur between the winter solstice (i.e., after December 20) and about February 20 of any year and show a distinctive peak 20–40 days after the winter solstice. Spawning typically occurred during periods of declining seawater temperature and slacking currents within a temperature range of 26–27.5°C and current speed of 2.5–3.5 cm s⁻¹. Males arrived early to spawning sites and stayed longer than females. These gender-based behavioral patterns are important to E. guttatus reproductive dynamics and must be factored into future studies and the design of fisheries regulations to ensure sustainability of spawning aggregation sites. The predictability of E. guttatus spawning aggregations relative to the winter solstice will be extremely beneficial for defining the temporal and spatial aspects of area closures. The consistency and synchrony of movement and migration will improve both the efficiency of planning research and monitoring programs and directing enforcement activities during critical time periods. Applying this knowledge strategically will maximize the limited resources available for research and enforcement and lead to greater protection of spawning aggregations.     

Reproduction in an aggregating grouper,the red hind,epinephelus guttatus

Synopsis We examined the reproductive pattern of an aggregating grouper, the red hind,Epinephelus guttatus, in Puerto Rico. Macroscopic and histological examination of gonads confirmed that, although mature, ripe ovaries are found over a three-month period, spawning activity is limited to about 2 weeks each year. Females are determinate spawners and individuals are able to spawn more than once during the course of the annual spawning season. The red hind is protogynous and 50% maturity is attained at 215 mm fork length. In western Puerto Rico, spawning occurs in aggregations at several sites within loosely defined areas located towards the edge of the insular platform. Sex ratios of individuals taken by hook and line at one of the aggregation areas over a consecutive six-year period, suggest considerable intra- and inter-annual variation most likely attributable to a combination of differential ingress and egress by males and females during the course of an aggregation and to fluctuations in recruitment, differential mortality by sex and sex change among years. A comparative assessment of the reproductive patterns of seven western AtlanticEpinephelus spp. suggests that aggregation-spawning is associated with medium- to large-sized groupers, while smaller groupers do not aggregate. Mating systems vary among congeners in association with trends in male-female size dimorphism, sexual pattern and sperm competition. The short-term, localized, nature of spawning aggregations renders species with this reproductive mode particularly vulnerable to fishing pressure at spawning sites. Aggregating species, therefore, may require special management consideration.     

Patagonian grouper Acanthistius patachonicus (Jenyns, 1840) forms massive spawning aggregations in the Southwest Atlantic

Here we describe massive spawning aggregations and seasonal changes in the large-scale distribution of the Patagonian grouper Acanthistius patachonicus in the Southwest Atlantic based on three sources of information: (a) data from bottom trawl research surveys covering the distributional range of the species within the Argentine continental shelf; (b) folk ecological knowledge gathered from experienced captains of the Argentine industrial trawl fisheries; and (c) sampling of an artisanal trap fishery targeting the Patagonian grouper in a specific location off the coast of Buenos Aires Province. The trawl surveys showed a general pattern of aggregation of Patagonian grouper towards the coast during the reproductive period (September–December). Captains described massive aggregations of the species at specific near-shore locations, where trawl catches of up to 15 t in a single haul were registered during the reproductive season. At a local scale, the artisanal trap fishery described operates exclusively during the reproductive period, targeting near-shore aggregations with a high proportion of individuals releasing gametes onboard. These three sources of information provide evidence of the existence of massive transient spawning aggregations of Patagonian grouper in the Argentine shelf. This is the first report of a reef fish spawning aggregation in the southern region of the Southwest Atlantic. Anecdotal information gathered in this study points to the depletion of many of the aggregations targeted during the 1980s and 1990s by the industrial fleet. At the same time, the spawning aggregation site off Buenos Aires Province has been productive for the last 34 years, being exploited exclusively by traps. The Patagonian grouper is classified as Data Deficient by the International Union for Conservation of Nature, which highlights the need for further research to determine its stock status. Mapping its current spawning aggregations should be a priority to inform the design of a targeted monitoring program and management plan for this species.     

Aggregation and spawning of the humphead wrasse Cheilinus undulatus (Pisces: Labridae): general aspects of spawning behaviour

The humphead wrasse Cheilinus undulatus formed resident spawning aggregations daily after high tide at specific locations along the seaward edge of the Palau barrier reef. The location and extent of one aggregation site remained consistent for 6 years with no physical features distinguishing it from adjacent areas. Spawning was documented most months and probably occurred year round with possible seasonal and lunar variation. Spawning males arrived first at the site, followed by females and potentially small primary males. The aggregation female to male sex ratio was estimated to be between 6:1 and 10:1. A maximum of 15 males and 100–150 females were observed at the site. A male courtship posture with the anal fin pointed, the caudal fin folded down and the dorsal fin folded against the body was maintained while swimming a few metres off the bottom in view of females. When ready to spawn females rose up as the posturing male passed and the pair released gametes in a relatively sedate fashion near the surface along the shelf break. No attempted predation on spawning adults was seen. Egg predation after spawning was uncommon. On days with early to mid‐day high tides the spawning period started 2·0–2·5 h after high tide when the speed of lagoon–ocean tidal currents peaked and lasted c. 1 h. On days with later afternoon high tides, spawning occurred sooner after high tide and before current speeds peaked. Other fishes with planktonic eggs spawned at the site as pairs or small groups in a rough succession after high tide with C. undulatus, the last species to spawn.     

Reproductive movement,residency and fisheries vulnerability of brown-marbled grouper, <Emphasis Type="Italic">Epinephelus fuscoguttatus</Emphasis> (Forsskål, 1775)

The brown-marbled grouper, Epinephelus fuscoguttatus, is a long-lived, late-maturing protogynous species listed as Near Threatened by the IUCN. In Pohnpei, Micronesia, reproductively active brown-marbled grouper were tagged with acoustic and spaghetti-type tags at a multi-species fish spawning aggregation (FSA) site to establish patterns of movement, residency and seasonality. Telemetry confirmed the use of common reproductive migratory corridors and significant sex-specific variations in residency at the FSA. Combined underwater visual census and telemetry data verified a 3-month peak aggregation period, with aggregations forming and persisting over ca. 12 days prior to full moon between January and May. FSA formation coincided with seasonally low and relatively stable seawater temperatures. Some males frequented the FSA site during each aggregation month over two consecutive years. Conversely, most females were present at the FSA during only a single aggregation period, with the month of visitation consistent among years. Nearly two-thirds of tagged fish were relocated or recaptured within 11 km of the aggregation site, with a maximum detected distance of 26 km and a minimum estimated catchment area of 100–175 km². Findings highlight the need for a combined approach to management that prohibits the capture and sale of reproductive adults and protects both spawning sites and common reproductive migratory corridors during aggregation periods.     

Reproduction of the Nassau grouper,Epinephelus striatus (Pisces: Serranidae) and its relationship to environmental conditions

Synopsis Spawning aggregations of the Nassau grouper, Epinephelus striatus, occurring at the southern end of Long Island were investigated during the winters of 1987–1988 and 1988–1989. Fish aggregate at two specific sites around the time of the full moon in December and January, possibly not during November and almost certainly not during February. At one site the fish aggregate well inside the shelf edge while at the other they are on the dropoff into deep water. Fish migrate, probably as distinct groups of up to about 500 individuals, to the sites on or before the full moon and remain for several days. There was no significant difference in the size of males and females caught from the aggregations between years or months. Female/male sex ratio was between 5 : 1 and 3 : 1. Courtship occurred in late afternoon with spawning commencing shortly before sunset. Most spawning occurred within 10 min of sunset. Water temperatures were 25.0–25.5°C during a period of gradual decrease towards the annual minimum in February and March. Two color patterns were important in courtship and spawning. The bicolor pattern is a submissive coloration indicating a non-aggressive state acquired by both males and females near the time of spawning. The dark phase is acquired by females who are followed by numerous bicolor fish during courtship and they lead spawning events in this pattern. Spawning occurred among subgroups of the aggregation numbering 3–25 fish; gamete release was well above the bottom. Drogues deployed with the gametes either moved inshore or did not move far away from the shelf edge over the course of several days. Nassau groupers may not strictly be protogynous hermaphrodites and other groupers which form spawning aggregations may not be so either. Latitudinal shifts in spawning time may be related to water temperatures. Currents at aggregation sites do not appear to favor offshore transport of eggs. Questions of whether spawning aggregations should be protected need to be answered.     

Spatial,temporal, and environmental dynamics of a multi-species epinephelid spawning aggregation in Pohnpei,Micronesia

Long-term and short-term underwater visual censuses using SCUBA, technical Nitrox, and closed circuit rebreathers (CCR) were carried out in Pohnpei, Micronesia, to define spatial and temporal dynamics within a semi-protected multi-species epinephelid (fish) spawning aggregation (FSA) of brown-marbled grouper, Epinephelus fuscoguttatus, camouflage grouper, Epinephelus polyphekadion, and squaretail coralgrouper, Plectropomus areolatus. Results identified species-specific patterns of habitat use, abundance, residency, and dispersal of FSAs. Fish spawning aggregations formed and dispersed monthly within a 21–160-d period after winter solstice within adjacent yet distinct outer reef habitats. The reproductive season coincided with periods of seasonally low sub-surface seawater temperature. Peaks in density varied among species both within the calendar year and relative to the winter solstice. Significant long-term declines in FSA density were observed for all three species, suggesting population-level fishery-induced impacts, similar to those previously reported for E. polyphekadion. Differences in density estimates were also observed between dive gear, with a threefold difference in densities measured by CCR for E. polyphekadion versus SCUBA that suggest a disturbance effect from exhaled SCUBA bubbles for this species. CCR also allowed surveys to be conducted over a larger area in a single dive, thereby improving the potential to gauge actual abundance and density within FSAs. Based on these findings, a combination of long-term and intensive short-term monitoring strategies is recommended to fully characterize trends in seasonal abundance and habitat use for aggregating species at single or multi-species FSA sites. Inherent variations in the timing and distribution of species within FSA make fine-scale temporal management protocols less effective than blanket protective coverage of these species at (e.g., marine protected areas covering FSAs and adjacent migratory corridors) and away from (i.e., temporal sales and catch restrictions) FSA sites.     

Variation in fish density,assemblage composition and relative rates of predation among mangrove,seagrass and coral reef habitats

We tested the hypothesis for several Caribbean reef fish species that there is no difference in nursery function among mangrove, seagrass and shallow reef habitat as measured by: (a) patterns of juvenile and adult density, (b) assemblage composition, and (c) relative predation rates. Results indicated that although some mangrove and seagrass sites showed characteristics of nursery habitats, this pattern was weak. While almost half of our mangrove and seagrass sites appeared to hold higher proportions of juvenile fish (all species pooled) than did reef sites, this pattern was significant in only two cases. In addition, only four of the six most abundant and commercially important species (Haemulon flavolineatum, Haemulon sciurus, Lutjanus apodus, Lutjanus mahogoni, Scarus iserti, and Sparisoma aurofrenatum) showed patterns of higher proportions of juvenile fish in mangrove and/or seagrass habitat(s) relative to coral reefs, and were limited to four of nine sites. Faunal similarity between reef and either mangrove or seagrass habitats was low, suggesting little, if any exchange between them. Finally, although relative risk of predation was lower in mangrove/seagrass than in reef habitats, variance in rates was substantial suggesting that not all mangrove/seagrass habitats function equivalently. Specifically, relative risk varied between morning and afternoon, and between sites of similar habitat, yet varied little, in some cases, between habitats (mangrove/seagrass vs. coral reefs). Consequently, our results caution against generalizations that all mangrove and seagrass habitats have nursery function.     

Forecasting lionfish sources and sinks in the Atlantic: are Gulf of Mexico reef fisheries at risk?

Invasive lionfish (Pterois volitans/miles complex) now permeate the entire tropical western Atlantic, Caribbean Sea, and Gulf of Mexico, but lionfish abundance has been measured only in select locations in the field. Despite its rapid range expansion, a comprehensive meta-population analysis of lionfish ‘sources’ and ‘sinks’ and consequentially the invader’s potential abundance and impacts on economically important, sympatric reef fishes have not been assessed. These data are urgently needed to spatially direct control efforts and to plan for and perhaps mitigate lionfish-caused damage. Here, we use a biophysical computer model to: (1) forecast larval lionfish sources and sinks that are also delineated as low to high lionfish ‘density zones’ throughout their invaded range, and (2) assess the potential vulnerability of five grouper and snapper species—Epinephelus morio, Mycteroperca microlepis, Epinephelus flavolimbatus, Lutjanus campechanus, and Rhomboplites aurorubens—to lionfish within these density zones in the Gulf of Mexico. Our results suggest that the west Florida shelf and nearshore waters of Texas, USA, and Guyana, South America, function both as lionfish sources and sinks and should be a high priority for targeted lionfish control. Furthermore, of the five groupers and snappers studied, the high fishery value E. morio (red grouper) is the Gulf of Mexico species most at risk from lionfish. Lacking a comprehensive lionfish control policy, these risk exposure data inform managers where removals should be focused and demonstrate the risk to five sympatric native groupers and snappers in the Gulf of Mexico that may be susceptible to dense lionfish aggregations, should control efforts fail.     

Seasonal occupancy and connectivity amongst nearshore flats and reef habitats by permit Trachinotus falcatus: considerations for fisheries management

We used acoustic telemetry to quantify permit Trachinotus falcatus habitat use and connectivity in proximity to the Florida Keys, USA, and assessed these patterns relative to current habitat and fisheries management practices. From March 2017 to June 2018, 45 permit tagged within 16 km of the lower Florida Keys were detected at stationary acoustic receivers throughout the south Florida region, the majority of which remained within the Special Permit Zone, where more extensive fisheries harvest regulations are implemented. There was a high level of connectivity between nearshore flats (i.e., <3 m water depth) and the Florida reef tract (FRT; 15–40 m water depth), with 75% of individuals detected in both habitats. These locations probably function primarily as foraging and spawning habitats, respectively. Permit occupancy on the FRT peaked during the months of March–September, with the highest number of individuals occurring there in April and May. Specific sites on the FRT were identified as potentially important spawning locations, as they attracted a high proportion of individuals that exhibited frequent visits with high residency durations. There were also significant positive relationships between seasonal habitat-use metrics on the FRT and an empirical permit gonadosomatic index. Large aggregations of permit at spawning sites on the FRT are potentially vulnerable to the effects of fishing (including predation during catch and release) at a critical point in their life cycle. These data on permit space use and movement, coupled with knowledge of stressors on their ecology, provide insights for implementing science-based strategic management plans.